Diversity of parasites of fish from the Upper Paraná River ... · zoa, three species of Phylum Sarcomastigophora were reported: Trypanosoma guairaensis Eiras, Rego and Pavanelli,

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Diversity of parasites of fish from the Upper Paraná River floodplain, Brazil

Takemoto, RM.*, Pavanelli, GC., Lizama, MAP., Lacerda, ACF., Yamada, FH., Moreira, LHA., Ceschini, TL. and Bellay, S.

Programa de Pós-graduação em Ecologia de Ambientes Aquáticos Continentais – PEA, Núcleo de Pesquisas em Limnologia, Ictiologia e Aquicultura – Nupélia,

Laboratório de Ictioparasitologia, Universidade Estadual de Maringá – UEM, Av. Colombo, 5790, CEP 87020-900, Maringá, PR, Brazil

*e-mail: [email protected]

Received November 27, 2008 – Accepted March 13, 2009 – Distributed June 30, 2009

(With 2 figures)

Abstract

The aim of the present study was to provide a current survey of the species of parasites found in the Upper Paraná River floodplain, as well as to investigate which strategies and mechanisms used by parasites, are favored and which environmental factors influence the parasite community in the studied environments. During a seven-year period from February 2000 to September 2007, 3,768 fish belonging to 72 species were collected and analyzed for the purpose of studying the parasite fauna. A total of 337 species of parasites were reported, including 12 new descrip-tions: one myxosporid, Henneguya paranaensis Eiras, Pavanelli and Takemoto, 2004; eight monogeneans, Kritskyia annakohnae Boeger, Tanaka and Pavanelli, 2001; Kritskyia boegeri Takemoto, Lizama and Pavanelli, 2002; Kritskyia eirasi Guidelli, Takemoto and Pavanelli, 2003; Demidospermus labrosi França, Isaac, Pavanelli and Takemoto, 2003; Demidospermus mandi França, Isaac, Pavanelli and Takemoto, 2003; Pseudovancleaveus paranaensis França, Isaac, Pavanelli and Takemoto, 2003; Tereancistrum curimba Lizama, Takemoto and Pavanelli, 2004 and Tereancistrum toksonum Lizama, Takemoto and Pavanelli, 2004; two digeneans, Sanguinicola platyrhynchi Guidelli, Isaac and Pavanelli, 2002 and Dadayius pacupeva Lacerda, Takemoto and Pavanelli, 2003 and one cestode, Nomimoscolex pertierrae Chambrier, Takemoto and Pavanelli, 2005. In addition, several other species were reported for the first time in new hosts or in the floodplain. Monogeneans presented the highest number of species, followed by digeneans. The infection site with the highest species richness was the intestine, with 127 species.

Keywords: ichthyoparasites, biodiversity, helminthes, crustaceans, parasite ecology.

Diversidade dos parasitos de peixes da planície de inundação do Alto Rio Paraná, Brasil

Resumo

O objetivo do presente trabalho foi realizar um novo levantamento das espécies de parasitos encontradas na planície de inundação do Alto Rio Paraná, bem como investigar quais estratégias e mecanismos utilizados pelos parasitos são mais favorecidos e quais fatores ambientais estão influenciando a comunidade parasitária nos ambientes estudados. Durante o período de fevereiro de 2000 a setembro de 2007, 3.768 espécimes de peixes pertencentes a 72 espécies fo-ram coletados e analisados, visando o estudo da fauna parasitária. Até o momento foram registradas, 337 espécies de parasitos, sendo 12 novas espécies: uma de mixosporídeo, Henneguya paranaensis Eiras, Pavanelli e Takemoto, 2004; oito de monogenéticos, Kritskyia annakohnae Boeger, Tanaka e Pavanelli, 2001; Kritskyia boegeri Takemoto, Lizama e Pavanelli, 2002; Kritskyia eirasi Guidelli, Takemoto e Pavanelli, 2003; Demidospermus labrosi França, Isaac, Pavanelli e Takemoto, 2003; Demidospermus mandi França, Isaac, Pavanelli e Takemoto, 2003; Pseudovancleaveus paranaensis França, Isaac, Pavanelli e Takemoto, 2003; Tereancistrum curimba Lizama, Takemoto e Pavanelli, 2004 e Tereancistrum toksonum Lizama, Takemoto e Pavanelli, 2004; duas de digenéticos, Sanguinicola platyrhynchi Guidelli, Isaac e Pavanelli, 2002 e Dadayius pacupeva Lacerda, Takemoto e Pavanelli, 2003 e uma de cestóide, Nomimoscolex pertierrae Chambrier, Takemoto e Pavanelli, 2005. Além dessas novas espécies, várias outras foram registradas pela primeira vez em novos hospedeiros ou na planície. O grupo dos monogenéticos foi o que apresentou maior número de espécies encontradas, seguido pelos digenéticos. O intestino foi o sítio de infecção que apresentou a maior riqueza, com 127 espécies.

Palavras-chave: ictioparasitos, biodiversidade, helmintos, crustáceos, ecologia de parasitos.

Takemoto, RM. et al.

692 Braz. J. Biol., 69(2, Suppl.): 691-705, 2009

1. Introduction

Floodplains are ecosystems that experience recurrent floods, favoring certain adaptations of the organisms in-habiting them and resulting in the presence of commu-nities with characteristic structures and functions (Junk, et al. 1989). Floods can alter the population dynamics of the ichthyofauna, as well as their physiological and bio-logical conditions, thereby influencing the structure and composition of the parasite fauna. In addition, chemical and physical variables of the water such as dissolved oxygen, temperature and turbulence can contribute to the emergence and increases in the demography of some parasite species (Pavanelli et al., 1997).

Luque and Poulin (2008) reported that differences in the taxonomic diversification of the parasite assem-blages of different fish species were mainly related to the environment (higher values in benthic-demersal spe-cies), trophic level (positive correlation) and tempera-ture (positive correlation for all groups of parasites in freshwater fishes). Therefore, the Upper Paraná River floodplain, characterized by a wide variety of habitats and species, favors the occurrence of a great diversity of fish parasites.

Lizama et al. (2006a) and Yamada et al. (2007) found higher parasitism indexes of monogeneans and digeneans in open lagoons in the Upper Paraná River floodplain. The authors explain that the presence of morphological adap-tations, such as eggs with adhesive appendices that allow for attachment to a substrate, could explain the higher number of monogeneans. Meanwhile, digeneans could be more prevalent in these environments due to the great quantity of intermediate hosts in open lagoons. Studying Pseudoplatystoma corruscans (Spix and Agassiz, 1829) and Schizodon borelli (Boulenger, 1900), Machado et al. (1995) observed that fish were more parasitized by ces-todes in semi-lentic environments and by nematodes in lentic environments, probably because of the great vari-ety of prey and availability of shelter.

The parasite fauna of fish from the Upper Paraná River floodplain was previously detailed by Pavanelli et al. (1997, 2004). The present study investigates which reproductive strategies and dispersion mechanisms are favored, and which environmental factors (internal and external) influence the parasite community. In addition, this paper complements previous studies presenting an updated list of species of parasites and hosts.

2. Materials and Methods

All fish were collected in the floodplain of the Upper Paraná River, Brazil, from February 2000 to September 2007. Fishes were captured using gill nets, and para-sites were collected with the aid of a stereoscopic mi-croscope. The eyes, skin, fins, gills, nostrils and mouth cavity of the selected individuals were studied for ec-toparasites. The body cavity was opened to examine the liver, stomach, pyloric caeca, intestine and gonads mi-croscopically for endoparasites (see Eiras et al., 2006).

The literature used to identify the parasites included original descriptions. Quantitative information on the prevalence and intensity of infection by different para-sites and on parasite community descriptors have been published previously for the following fish species: Prochilodus lineatus (Valenciennes, 1837) (Takemoto et al., 2002; Eiras et al., 2004; Lizama et al., 2004; 2005; 2006a, b); Acestrorhynchus lacustris (Lütken, 1875) (Carvalho et al., 2003); Iheringichthys labrosus (Lütken, 1874) (França et al., 2003); Hemisorubim platyrhynchos (Valenciennes, 1840) (Guidelli et al., 2003a); Leporinus spp. (Guidelli et al., 2003b; 2006); Metynnis maculatus (Kner, 1858) (= M. lippincottianus) (Lacerda et al., 2003); A. lacustris, S. borellii, P. lineatus, Serrasalmus marginatus Valenciennes, 1837 and S. maculatus Kner, 1858 (Lacerda et al., 2007); Potamotrygon spp. (Lacerda et al., 2008; 2009); Hoplias aff. malabaricus (Bloch, 1794), Satanoperca pappaterra (Heckel, 1840), Plagioscion squamosissimus (Heckel, 1840), Crenicichla britskii Kullander, 1982, Cichla monoculus Spix and Agassiz, 1831 (= C. kelberi) and Cichlasoma paranaense Kullander, 1983 (Aoyama et al., 2005); P. squamosissimus (Tavernari et al., 2005); P. corruscans (Chambrier et al., 2006); Loricariichthys platymetopon Isbrücker and Nijssen, 1979 (Ferrari et al., 2007); S. papaterra (Yamada et al., 2007) and Cichla spp. (Yamada et al., 2008).

3. Results

We collected and analyzed 3,768 fish belonging to 72 species. A total of 337 species of parasites were re-ported (Appendix 1), including the description of 12 new species: one myxosporid, Henneguya paranaensis Eiras, Pavanelli and Takemoto, 2004; eight monogeneans, Kritskyia annakohnae Boeger, Tanaka and Pavanelli, 2001; K. boegeri Takemoto, Lizama and Pavanelli, 2002; K. eirasi Guidelli, Takemoto and Pavanelli, 2003; Demidospermus labrosi França, Isaac, Pavanelli and Takemoto, 2003; D. mandi França, Isaac, Pavanelli and Takemoto, 2003; Pseudovancleaveus paranaensis França, Isaac, Pavanelli and Takemoto, 2003; Tereancistrum curimba Lizama, Takemoto and Pavanelli, 2004 e T. toksonum Lizama, Takemoto and Pavanelli, 2004; two digeneans, Sanguinicola platyrhynchi Guidelli, Isaac and Pavanelli, 2002 and Dadayius pacupeva Lacerda, Takemoto and Pavanelli, 2003 and one cestode, Nomimoscolex pertierrae Chambrier, Takemoto and Pavanelli, 2006.

In total, 12 groups of parasites were collected, mainly Platyhelminthes (Monogenea, Digenea and Cestoda) and Nematoda. Among the groups of parasites found, mono-geneans presented the greatest number of species with 82 (Figure 1), including species that parasitized gills, nasal fossae and urinary bladder. Among digeneans and nema-todes, which also presented high numbers of species, adults and larval forms were observed. Sixty-nine species of digeneans were recorded, with 28 reported as larvae.

Diversity of fish parasites

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Figure 1. Species richness of fish parasites according to zoological group reported in the Upper Paraná River flood-plain.

Figure 2. Species richness according to the site of infection/infestation reported in the Upper Paraná River floodplain.

Nine out of 66 species of nematodes were larvae, and Contracaecum sp. (Anisakidae) was the most common.

The main infection site was the intestine, which was parasitized by 127 different species of parasites, including digeneans, cestodes, nematodes and acanthocephalans. These species were always found as adults (Figure 2). The gills presented the second highest number of species (91), mainly monogeneans and copepods. In the mesen-tery, the helminthes were mainly larval forms.

The parasitic protozoa are usually classified accord-ing to their motility mechanisms. These include amoe-boid, flagellated, and ciliated forms. Among the proto-zoa, three species of Phylum Sarcomastigophora were reported: Trypanosoma guairaensis Eiras, Rego and Pavanelli, 1989, T. scrofae Eiras and Pavanelli, 1989 and T. nupelianus Eiras, Rego and Pavanelli, 1990, parasitizing Megalancistrus parananus (Peters, 1881), P. lineatus and Rhineleps aspera Spix and Agassiz, 1829, respectively.

The myxozoans, which until recently were consid-ered protozoans, are now considered metazoans. The genus Henneguya was reported in the floodplain, with the description of a new species, H. paranaensis Eiras, Pavanelli and Takemoto, 2004.

During this period, the class Monogenea had the largest number of new species.

Digeneans presented the second highest number of species and low host specificity. Some species that parasitize fish in the larval stage, such as Diplostomum (Austrodiplostomum) compactum (Lutz, 1928) (prob-ably introduced), were reported in nine different spe-cies of hosts: H. aff. malabaricus, S. pappaterra, P. squamosissimus, C. britskii, Cichla kelberi Kullander and Ferreira, 2006 (= Cichla monoculus), C. paranaense, Hypostomus regani (Ihering, 1905), S. borellii, and Auchenipterus osteomystax (Miranda Ribeiro, 1918).

Nematodes also generally have low host specificity, such as the nematode Procamallanus (Spirocamallanus)

inopinatus Travassos, Artigas and Pereira, 1928, which was reported in nine different species of hosts (S. marginatus, S. maculatus, S. borelli, H. aff. malabaricus, M. lippincottianus, Leporinus elongatus Valenciennes, 1850, L. obtusidens (Valenciennes, 1837), L. lacustris Amaral Campos, 1945 and P. corruscans) in the current study. However, Moravec (1998) had al-ready registered this species in more than 40 species of fish from the Neotropical region. Another species of nematode that presented low specificity was the larvae of Contracaecum sp., which was already reported in 17 dif-ferent species of hosts. This species deserves special at-tention because it parasitizes fish as larvae, using it as an intermediate host and then presenting zoonotic potential.

During this period only one species of cestode was de-scribed, Nomimoscolex pertierrae Chambrier, Takemoto and Pavanelli, 2006. In previous studies, the occurrence of this group was observed mainly in Siluriform fishes and with high host specificity. Considering only the fish “pintado”, P. corruscans and “jaú”, Zungaro zungaro (Humboldt, 1821) (= Paulicea luetkeni), six species of proteocephalideans were reported in each species, all adults. Larval stages of Cyclophyllidea were also report-ed, including Valipora campylancristrota (Wedl, 1855) in P. lineatus, Pimelodus maculatus Lacépède, 1803 and Hoplosternum littorale (Hancock, 1828), all para-sitizing the gall bladder. In the stingrays Potamotrygon motoro (Müller and Henle, 1841) and P. falkneri Castex and Maciel, 1963, cestodes of orders Tetraphyllidea, Acanthobothrium regoi Mayes, Brooks and Thorson, 1981, Rhinebothrium paratrygoni Rego and Dias, 1976, Potamotrygonocestus travassosi Rego, 1979 and Trypanorhyncha, Paroncomegas araya (Woodland, 1934) were reported.

Among the acanthocephalans, Quadrigyrus machadoi Fabio, 1983 presented low specificity, oc-curring in seven different species of hosts (P. motoro,


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P. falkneri, H. aff. malabaricus, Gymnotus carapo Linnaeus, 1758, P. maculatus, H. platyrhynchos and C. kelberi). Acanthocephalans of the genus Quadrigyrus occurred as adults and larvae.

The main families of Copepoda reported in the flood-plain were Ergasilidae and Vaigamidae, parasitizing the branchial arcs and the nostrils, respectively. In addition, copepods of family Therodamasidae were observed par-asitizing the branchial arcs of P. maculatus. This family is characterized by an anterior structure that penetrates into the tissues of the branchial arcs of the host, causing considerable lesions. This is the first record of this fam-ily in the Upper Paraná River floodplain.

Branchiura, Isopoda, Hirudinea, Pentastomida and Acarina presented low species diversity.

4. Discussion

In previous studies, Takemoto et al. (2004) reported 184 species of fish parasites in the Upper Paraná River floodplain. Currently, 337 species are known. This aug-mentation of species richness is due to the increase in the quantity of studies conducted in recent years, which have included different species of fish.

In addition to the previously described species, several other species were reported for the first time in the floodplain. In P. maculatus, for example, we reported the first copepods of the family Therodamasidae, and this is also the first record for the Upper Paraná River basin.

Myxozoa is a group of parasitic organisms for which the basic knowledge regarding phylogenetic affinities and life cycles has changed dramatically in the last two decades. The myxozoans were included as unicellular protozoans. However, according to Smothers et al. (1994) and Siddall et al. (1995), recent evidence clearly indi-cates that myxozoans are true metazoans. In the Upper Paraná River floodplain, only the genus Henneguya was observed.

Monogenea is the group that has presented the great-est number of species so far. Monogeneans are a diverse group of parasites that exhibit a relatively high degree of host specificity when compared to other groups of para-sites. Because monogeneans do not represent a zoonotic problem for humans, and because most of them are not pathogenic, very little is known about the biology of most species (Bush et al., 2001).

The number of species recorded in this class tends to increase in the floodplain, since previous studies have already shown its high species richness in every species of host.

Among the groups of parasites found, digeneans pre-sented the second highest number of species (69), includ-ing adults and larval stages. Some species of digeneans presented low specificity and were mainly present in lar-val forms. As an example we can cite D. (A.) compactum, a digenean that, in the metacercaria stage, parasitizes the eyes of fish. This parasite was probably introduced to the floodplain with P. squamosissimus. Despite being almost

an obligatory parasite of this species, we noticed that in recent years, the number of species of parasitized hosts has increased considerably. We recorded this digenean in nine species of fish, showing low specificity and rapid adaptation to the new environment.

The conservation of the native fauna depends on re-search to increase knowledge regarding which parasites spread globally (Font, 2003). The goal of ecologists is not only to document the distribution of parasites, but also to determine methods by which parasites can disperse to new areas. Through the determination of the groups of parasites that can establish themselves in a new environ-ment, it is possible to determine which strategies of re-production are favored. Improved understanding of these mechanisms of dispersion can increase the chances of limiting the dispersion of certain parasites (Font, 2003).

Digeneans represent the highest number of species because fish can act as intermediate hosts, harboring lar-val stages, and also as definitive hosts, harboring adult forms. Moreover, the majorities of the fish examined here occupy intermediate levels in the food chain and can be parasitized by both forms.

Another larvae that showed low specificity was the nematode Contracaecum sp. This species belongs to the family Anisakidae and is of great importance in public health because of its high zoonotic potential. Anisakids are important to humans on a number of counts: econom-ically, politically and mainly pathologically. Paratenic hosts are often fish that are important for human con-sumption. The worms do not mature in humans and the pathology is mostly associated with penetration of the small intestine and stomach by the larvae. The degree of discomfort is likely a consequence of the number of larvae penetrating. In rare cases, humans have died from anisakiasis due to severe peritonitis (Bush et al., 2001). However, there are no records of this zoonosis in the re-gion and this is probably due to the fact that all of the lar-vae of this species were encysted in the mesentery, which is not used for human consumption.

The occurrence of a few species of some groups may be due to the absence or low number of certain hosts required for a particular stage of the developmental cy-cle. This is probably the case for the pentastomids that parasitize fish as larvae and alligators as adults. Although there are alligators in the floodplain, their numbers are relatively small when compared to other regions. The pentastomids are a small group of parasites that includes about 100 species. Studies on the phylogenetic relation-ships of the Pentastomida, a group traditionally treated as a separate phylum, suggest that they are truly arthro-pods with possible affinities for the crustacean (Bush et al., 2001).

The diversity of endohelminths can vary according to the environment or the species of host, and can be related to the size, longevity and particularly the food of the host. The intermediate host is necessarily part of the diet of the definitive host (Luque and Poulin, 2008). Furthermore, fish with a longer lifespan present more


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time of exposure to parasites, favoring cumulative proc-esses in the host.

In the Upper Paraná River floodplain, Takemoto et al. (2005) studied the effects of several host traits (body size, social behavior, reproductive behavior, spawning type, trophic category, feeding habits, relative position in the food web, preference for certain habitats and whether the fish species are native or exotic) on metazoan en-doparasite species richness. The CPUE (number of in-dividuals of a given fish species captured per 1,000 m2 of net during 24 hours) was the sole significant predictor of parasite species richness, regardless of control for the confounding influences of host phylogeny and sampling effort. The results suggest that in the Upper Paraná River floodplain (with homogeneous physical characteristics and occurrence of many flood pulses), population den-sity of different host species might be the major deter-minant of the parasite species richness. This result is in concordance with epidemiological theory predicting that hosts with larger or denser populations will more readily sustain several parasite populations. Because parasites in this study use fish as either intermediate or definitive hosts, depending on species, this situation might reflect a combination of large intermediate and definitive host populations facilitating parasite transmission and repro-duction, respectively. However, it is important to mention that the cited study was developed utilizing data obtained from 1986 to 2003, and current floods pulses do not oc-cur periodically as they used to because of the construc-tion of several hydroelectric power plants upstream from the studied area.

To date, 182 species of fish have been registered in the Upper Paraná River floodplain (Graça and Pavanelli, 2007). If we assume that each species of host can harbor an average of ten species of parasites (Takemoto et al., 2004), there should be about 1,820 species of parasites in the floodplain. However, in the present study we re-corded less than 350 species of parasites, with many more to be studied. This number, however, may be much higher, considering that in some hosts such as P. lineatus, 33 species of parasites were reported.

To understand the role of the community of parasites in an ecosystem, previous knowledge of the species that compose it is required. Continuation of studies using taxonomic and systematic approaches is the key to un-derstanding how biotic and abiotic factors affect species, since there is no way to understand the effects on a popu-lation without knowing the species.

Acknowledgements — We would like to express our gratitude to several people who helped us to obtain the data presented in this work: Gislaine M Guidelli, Patrícia M Machado Aoyama, Andréia Isaac, Solange de Carvalho, Ana Paula Ferrari, Jakeline G França, Fernando de C Tavernari, Kennya F Ito, Sara T Moreira, Eliane da S Fernandes, Luciana MS Hino, Letícia C Karling, Filipe MS Alvarenga, Paula G Milani and Ana Cláudia Munhoz RM Takemoto and GC Pavanelli were supported by a Research Fellowship from CNPq (Conselho Nacional de Pesquisa e Desenvolvimento Tecnológico, Brazil). ACF Lacerda, FH Yamada and S Bellay were supported by Masters

Scholarships from CAPES (Coordenação de Aperfeiçoamento de Pessoal de Nível Superior). LHA Moreira was also supported by CNPq (Master Scholarship).

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______, 2006a. Influence of the seasonal and environmental patterns and host reproduction on the metazoan parasites of Prochilodus lineatus (Valenciennes, 1836) (Prochilodontidae) of the Upper Paraná River floodplain, Brazil. Brazilian Archives of Biology and Technology, vol. 49, no. 4, p. 611-622.

______, 2006b. Parasitism influence on the Hepato, Splenosomatic and Weight/Length relation and Relative Condition Factor of Prochilodus lineatus (Valenciennes, 1836) (Prochilodontidae) of the Upper Paraná River Floodplain, Brazil. Revista Brasileira de Parasitologia Veterinária, vol. 15, no. 3, p. 116-122.

LUQUE, JL. and POULIN, R., 2008. Linking ecology with parasite diversity in Neotropical fishes. Journal of Fish Biology, vol. 72, no. 1, p. 189-204.

MACHADO, MH., PAVANELLI, GC. and TAKEMOTO, RM., 1995. Influence of the type of environment and the

hidrological level variation in the endoparasitic infrapopulations of Pseudoplatystoma corruscans and Schizodon borelli (Osteichthyes) of the high river Paraná. Revista Brasileira de Zoologia, vol. 12, no. 4, p. 961-976.

MORAVEC, F., 1998. Nematodes of freshwater fishes of the neotropical region. Praha: Academia. 464p.

PAVANELLI, GC., MACHADO, MH., TAKEMOTO, RM., 1997. Fauna helmíntica de peixes do rio Paraná, região de Porto Rico, Paraná. In VAZZOLER, AEAM., AGOSTINHO, AA. and HAHN, NS. (Eds.). A Planície de inundação do Alto rio Paraná: aspectos físicos, biológicos e sócioeconômicos. Maringá: Eduem. p. 307-329.

PAVANELLI, GC., MACHADO, MH., TAKEMOTO, RM., GUIDELLI, GM. and LIZAMA, MAP., 2004. Helminth fauna of fishes: diversity and ecological aspects. In THOMAZ, SM., AGOSTINHO, AA. and HAHN, NS. (Eds.). The Upper Paraná river and its Floodplain: physical aspects, ecology and conservation. Leiden: Backhuys Publishers. p. 309-329.

SIDDALL, ME., MATIN, DS., BRIDGE, D., DESSER, SS. and CONE, DL., 1995. The demise of a phylum protists: phylogeny of Myxozoa and other parasitic Cnidaria. Journal of Parasitology, vol. 81, no. 6, p. 961-967.

SMOTHERS, JF., Von DOHLEN, CD., SMITH Jr., LH. and SPALL, RD., 1994. Molecular evidence that myxozoan protists are metazoans. Science, vol. 265, no. 5179, p. 1719-1721.

TAKEMOTO, RM., LIZAMA, MAP. and PAVANELLI, GC., 2002. A new species of Kritskyia (Dactylogyridae, Ancyrocephalinae) parasite of urinary bladder of Prochilodus lineatus (Prochilodontidae, Characiformes) from the floodplain of the high Paraná river, Brazil. Memórias do Instituto Oswaldo Cruz, vol. 97, no. 3, p. 313-315.

TAKEMOTO, RM., LIZAMA, MAP., GUIDELLI, GM. and PAVANELLI, GC., 2004. Parasitos de peixes de águas continentais. In RANZANI-PAIVA, MJT., TAKEMOTO, RM. and LIZAMA, MAP. (Eds.). Sanidade de organismos aquáticos. São Paulo: Editora Varela. p. 179-197.

TAKEMOTO, RM., PAVANELLI, GC., LIZAMA, MAP., LUQUE, JL. and POULIN, R., 2005. Host density as a major determinant of endoparasite species richness in fishes of floodplain of the upper Parana River, Brazil. Journal of Helminthology, vol. 79, no. 1, p. 75-84.

TAVERNARI, FC., BELLAY, S., TAKEMOTO, RM., GUIDELLI, GM., LIZAMA, MAP. and PAVANELLI, GC., 2005. Ecological aspects of Diplectanum piscinarius (Platyhelminthes, Monogenea) parasite of gills of Plagioscion squamosissimus (Osteichthyes, Sciaenidae) in the Upper Paraná River floodplain, Brazil. Acta Scientiarum, vol. 27, no. 3, p. 225-229.

YAMADA, FH., TAKEMOTO, RM. and PAVANELLI, GC., 2007. Ecological aspects of ectoparasites from gills of Satanoperca pappaterra (Heckel, 1840) (Cichlidae) from the Upper Paraná River floodplain, Brazil. Acta Scientiarum, vol. 29, no. 3, p. 331-336.

______, 2008. Relação entre fator de condição relativo (Kn) e abundância de ectoparasitos de brânquias em duas espécies de ciclídeos da bacia do rio Paraná, Brasil. Acta Scientiarum, vol. 30, no. 2, p. 213-217.


697Braz. J. Biol., 69(2, Suppl.): 691-705, 2009

Appendix 1. Continued...Appendix 1. Species list of hosts (arranged by family) and parasites reported by Pavanelli et al. (1997, 2004) and the present work in the Upper Paraná River floodplain.

Species listCHONDRICHTHYESMYLIOBATIFORMES

Potamotrygonidae• Potamotrygon motoro (Müller and Henle, 1841) (raia)

CestodaAcanthobothrium regoi Mayes, Brooks and Thorson, 1981Rhinebothrium paratrygoni Rego and Dias, 1976

AcanthocephalaQuadrigyrus machadoi Fabio, 1983

• Potamotrygon falkneri Castex and Maciel, 1963 (raia)

DigeneaClinostomum complanatum Rudolphi, 1814 (metacercariae)Tylodelphis sp. (metacercariae)Genarchella sp.

CestodaAcanthobothrium regoi Mayes, Brooks and Thorson, 1981Rhinebothrium paratrygoni Rego and Dias, 1976Potamotrygonocestus travassosi Rego, 1979Paroncomegas araya (Woodland, 1934)

NematodaBrevimulticaecum sp. (larvae) Cucullanus sp.Echinocephalus sp.Spinitectus sp.

AcanthocephalaQuadrigyrus machadoi Fabio, 1983

OSTEICHTHYESCHARACIFORMES

Characidae• Aphyocharax anisitsi Eigenmann and Kennedy, 1903 (piqui)

MonogeneaDigenea

DiplostomidaeCestoda

Proteocephalidea (plerocercoid)NematodaAcanthocephala

Quadrigyrus sp.

• Psellogrammus kennedyi (Eigenmann, 1903) (lambari)

DigeneaDiplostomidae

CestodaProteocephalidea (plerocercoid)

Nematoda

Species listContracaecum sp. (larvae)

AcanthocephalaQuadrigyrus sp. (larvae)

• Serrapinus notomelas (Eigenmann, 1915) (pequira)Cysts

• Astyanax altiparanae Garutti and Britski, 2000 (tambiú)

MonogeneaUrocleidoides sp.Amphithecium sp.

DigeneaClinostomum sp. (metacercariae)Herpetodiplostomum sp. (metacercariae)Tylodelphys sp. (metacercariae)Ascocotyle sp. (metacercariae)Antorchis sp. (metacercariae)Bucephalidae (metacercariae)


NematodaProcamallanus (Spirocamallanus) caballeroi Bashirullah, 1977Contracaecum sp. (larvae)Spiroxys sp.

AcanthocephalaQuadrigyrus sp.

CopepodaBrasergasilus sp.Ergasilus sp.Vaigamus sp.Acusicola sp.

• Moenkhausia intermedia Eigenmann, 1908 (lambari)

DigeneaCopepoda

• Moenkhausia sanctaefilomenae (Steindachner, 1907) (lambari-do-olho-vermelho)

MonogeneaJainus hexops Kritsky and Leiby, 1972

• Roeboides descalvadensis Fowler, 1932 (dentudo)Digenea (metacercariae)Monogenea

Jainus sp.Demidospermus sp.


Copepoda

• Salminus brasiliensis (Cuvier, 1816) (dourado)MyxosporeaDigenea

Neocladocystis intestinalis (Vaz,1932)Prosthenhystera obesa (Diesing, 1850)

Monogenea


698 Braz. J. Biol., 69(2, Suppl.): 691-705, 2009

Appendix 1. Continued... Appendix 1. Continued...

Species listJainus sp.Anacanthorus sp.Rhinoxenus bulbovaginatus Boeger, Domingues and Pavanelli, 1995

CestodaMonticellia coryphicephala (Monticelli, 1891)

NematodaEustrongylides ignotus Jäegerskiold, 1909 (larvae)Anisakidae

BranchiuraDolops longicauda (Heller, 1857)Dolops sp.

• Acestrorhynchus lacustris (Lütken, 1875) (peixe-cachorro)

MonogeneaDigenea

Rhipidocotyle gibsoni Kohn and Fernandes, 1994Clinostomum sp. (metacercariae)

CestodaNematoda

Contracaecum sp. (larvae)Procamallanus sp. (larvae)Eustrongylides sp. (larvae)Philometridae

AcanthocephalaQuadrigyrus torquatus Van Cleave, 1920

CopepodaRhinergasilus piranhus Boeger and Thatcher, 1988

• Galeocharax knerii (Steindachner, 1879) (peixe-cadela)

CestodaProteocephalidea

Isopoda

• Brycon orbignyanus (Valenciennes, 1850) (piracanjuba)

Nematoda (larvae)

Serrasalmidae• Serrasalmus marginatus Valenciennes, 1837 (piranha)

MonogeneaKritskyia annakohnae Boeger, Tanaka and Pavanelli, 2001Rhinoxenus sp.Anacanthorus sp.1Anacanthorus sp.2Anacanthorus sp.3Amphithecium sp.

DigeneaAscocotyle sp. (metacercariae)

NematodaProcamallanus (Spirocamallanus) inopinatus Travassos, Artigas and Pereira, 1928

Species listCucullanus sp.Eustrongylides ignotus Jäegerskiold, 1909 (larvae)PhilometridaeCapilariidae

AcanthocephalaCopepoda

Gamispatulus schizodontis Thatcher and Boeger, 1984

Acarina

• Serrasalmus maculatus Kner, 1858 (piranha)Monogenea

Kritskyia annakohnae Boeger, Tanaka and Pavanelli, 2001

DigeneaProsorhynchus piranhus Thatcher, 1999

CestodaProteocephalus serrasalmus Rego and Pavanelli, 1990

NematodaProcamallanus (Spirocamallanus) inopinatus Travassos, Artigas and Pereira, 1928Cucullanus sp.Eustrongylides ignotus Jäegerskiold, 1909 (larvae)PhilometridaeCapilariidae

AcanthocephalaCopepoda

Gamispatulus schizodontis Thatcher and Boeger, 1984

• Myloplus cf. tiete Eigenmann and Norris, 1900 (pacu-prata)

Digenea

• Metynnis lippincottianus (Cope, 1870) (pacu-peva)Digenea

Dadayius pacupeva Lacerda, Takemoto and Pavanelli, 2003

NematodaProcamallanus (Spirocamallanus) inopinatus Travassos, Artigas and Pereira, 1928Spinoxyuris oxydoras Petter, 1994Raphidascaris (Sprentascaris) mahnerti (Petter and Cassone, 1984)Contracaecum sp.

• Piaractus mesopotamicus (Holmberg, 1887) (pacu)Digenea

Dadaytrema oxycephala (Diesing, 1836)Cestoda

Proteocephalus vazollerae Pavanelli and Takemoto, 1995

NematodaRondonia rondoni Travassos, 1920

Acanthocephala


699Braz. J. Biol., 69(2, Suppl.): 691-705, 2009


Species list

Anostomidae• Leporellus vittatus (Valenciennes, 1850) (solteira)

MonogeneaJainus sp.

DigeneaCreptotrema creptotrema Travassos, Artigas and Pereira, 1928


NematodaContracaecum sp.

• Leporinus macrocephalus Garavello and Britski, 1988 (piaussú)

MonogeneaRhinoxenus sp.

Digenea (metacercariae)

• Leporinus friderici (Bloch, 1794) (piau)Monogenea

Cleidodiscus sp.Tereancistrum parvus Kritsky, Thatcher and Kayton, 1980Tereancistrum sp.Urocleidoides paradoxus Kritsky, Thatcher and Boeger, 1986Urocleidoides sp.Jainus sp.Kritskyia eirasi Guidelli, Takemoto and Pavanelli, 2003Rhinoxenus arietinus Kritsky, Boeger and Thatcher, 1988

DigeneaCreptotrema creptotrema Travassos, Artigas and Pereira, 1928Diplostomum sp. (metacercariae)Echinostomatidae (metacercariae)Herpetodiplostomum sp. (metacercariae)Paralecithobotrys brasiliensis Freitas, 1947Saccocoelioides saccodontis Thatcher, 1978

CestodaProteocephalus vazzolerae Pavanelli and Takemoto, 1995

NematodaBrevimulticaecum sp. (larvae)Contracaecum sp. (larvae)Cystidicoloides sp. (larvae)Dycheline leporini Petter, 1989Goezia brevicaeca Moravec, Kohn and Fernandes, 1994Goezia spinulosa (Diesing, 1839)Hysterothylacium sp. (larvae)Procamallanus (Spirocamallanus) amarali Vaz and Pereira, 1934Procamallanus (Spirocamallanus) iheringi Travassos, Artigas and Pereira, 1928Procamallanus (Spirocamallanus) inopinatus Travassos, Artigas and Pereira, 1928

Species listBranchiura

Argulus sp.Dolops nana Lemos de Castro, 1950

CopepodaGamispatulus schizodontis Thatcher and Boeger, 1984Vaigamidae

• Leporinus elongatus Valenciennes, 1850 (piau)Monogenea

Tereancistrum parvus Kritsky, Thatcher and Kayton, 1980Tereancistrum sp.Urocleidoides paradoxus Kritsky, Thatcher and Boeger 1986Urocleidoides sp.Jainus sp.Kritskyia sp.Rhinoxenus arietinus Kritsky, Boeger and Thatcher, 1988

DigeneaCreptotrema creptotrema Travassos, Artigas and Pereira, 1928Herpetodiplostomum sp. (metacercariae)Paralecithobotrys brasiliensis Freitas, 1947Saccocoelioides magniovatus Szidat, 1954

NematodaBrevimulticaecum sp. (larvae)Capillostrongyloides sentinosa (Travassos, 1927)Contracaecum sp. (larvae)Goezia spinulosa (Diesing, 1839)Porrocaecum sp. (larvae)Procamallanus (Spirocamallanus) amarali Vaz and Pereira, 1934Procamallanus (Spirocamallanus) iheringi Travassos, Artigas and Pereira, 1928Procamallanus (Spirocamallanus) inopinatus Travassos, Artigas and Pereira, 1928

BranchiuraDolops nana Lemos de Castro, 1950Dolops sp.

CopepodaErgasilus bryconis Thatcher, 1981Gamispatulus schizodontis Thatcher and Boeger, 1984Vaigamidae

• Leporinus obtusidens (Valenciennes, 1837) (piapara)

MonogeneaCleidodiscus sp.Tereancistrum parvus Kritsky, Thatcher and Kayton, 1980Tereancistrum sp.Urocleidoides paradoxus Kritsky, Thatcher and Boeger, 1986Urocleidoides sp.


700 Braz. J. Biol., 69(2, Suppl.): 691-705, 2009


Species listJainus sp.Kritskyia sp.Rhinoxenus arietinus Kritsky, Boeger and Thatcher, 1988

DigeneaCreptotrema lynchi Brooks, 1976Herpetodiplostomum sp. (metacercariae)Megacoelium sp.Neodiplostomum sp. (metacercariae)Paralecithobotrys brasiliensis Freitas, 1947

NematodaBrevimulticaecum sp. (larvae)Contracaecum sp. (larvae)Eustrongylides sp. (larvae)Goezia spinulosa (Diesing, 1839)Procamallanus (Spirocamallanus) amarali Vaz and Pereira, 1934Procamallanus (Spirocamallanus) iheringi Travassos, Artigas and Pereira, 1928Procamallanus (Spirocamallanus) inopinatus Travassos, Artigas and Pereira, 1928

BranchiuraArgulus sp.Dolops nana Lemos de Castro, 1950Dolops sp.

CopepodaErgasilus bryconis Thatcher, 1981Gamispatulus schizodontis Thatcher and Boeger, 1984Amplexibranchius sp.Vaigamidae

• Leporinus lacustris Amaral Campos, 1945 (corró)Monogenea

Cleidodiscus sp.Tereancistrum parvus Kritsky, Thatcher and Kayton, 1980Tereancistrum sp.Urocleidoides paradoxus Kritsky, Thatcher and Boeger, 1986Urocleidoides sp.Jainus sp.Kritskyia eirasi Guidelli, Takemoto and Pavanelli, 2003Rhinoxenus arietinus Kritsky, Boeger and Thatcher, 1988

DigeneaChalcinotrema thatcheri Kohn, Fernandes and Gibson, 1999Clinostomum complanatum Rudolphi, 1814 (metacercariae)Creptotrema sp.Cystodiplostomum sp.(metacercariae)Herpetodiplostomum sp. (metacercariae)Saccocoelioides magnus Szidat, 1954Tylodelphis sp. (metacercariae)

Species listCestoda

Proteocephalus vazzolerae Pavanelli and Takemoto, 1995

NematodaAncyracanthus schubarti Kohn, Gomes and Motta, 1968Brevimulticaecum sp. (larvae)Contracaecum sp. (larvae)Dycheline leporini Petter, 1989Procamallanus (Spirocamallanus) inopinatus Travassos, Artigas and Pereira, 1928

AcanthocephalaOctospiniferoides incognita Schmidt and Hugghins, 1973Quadrigyrus torquatus Van Cleave, 1920

BranchiuraArgulus sp.

CopepodaErgasilus bryconis Thatcher, 1981Gamispatulus schizodontis Thatcher and Boeger, 1984Vaigamidae

• Schizodon borelli (Boulenger, 1900) (piava)Digenea

Ithyoclinostomum dimorphum (Diesing, 1850)Clinostomum sp. (metacercariae)Saccocoelioides platensis Lunaschi, 1984Paralecithobotrys brasiliensis Freitas, 1947Diplostomum (Austrodiplostomum) compactum (Lutz, 1928)

MonogeneaJainus sp.Urocleidoides sp.Tereancistrum sp.

NematodaProcamallanus (Spirocamallanus) inopinatus Travassos, Artigas and Pereira, 1928Procamallanus (Spirocamallanus) iheringi Travassos, Artigas and Pereira, 1928Cucullanus pinnai Travassos, Artigas and Pereira, 1928Ancyracanthus schubarti Kohn, Gomes and Motta, 1968

AcanthocephalaOctospiniferoides incognita Schmidt and Hugghins, 1973Echinorhynchus sp.

CopepodaGamispatulus schizodontis Thatcher and Boeger, 1984

BranchiuraDolops sp.

• Schizodon altoparanae Garavello and Britski, 1990 (piava)

Nematoda


701Braz. J. Biol., 69(2, Suppl.): 691-705, 2009


Species listProcamallanus sp.

Acanthocephala

• Schizodon nasutus Kner, 1858 (piava)Monogenea

Curimatidae• Cyphocharax nagelii (Steindachner, 1881) (saguiru)

DigeneaDiplostomum sp. (metacercariae)Tylodelphis sp. (metacercariae)

• Cyphocharax modestus (Fernández-Yépez, 1948) (saguiru)

Digenea (metacercariae)

• Steindachnerina insculpta (Fernández-Yépez, 1948) (saguiru)

MonogeneaUrocleidoides sp.

DigeneaDiplostomum sp. (metacercariae)

NematodaTravnema travnema Pereira,1938Cosmoxynema vianai Travassos, 1949

Prochilodontidae• Prochilodus lineatus (Valenciennes, 1837) (curimba)

ProtozoaTrypanosoma scrofae Eiras and Pavanelli, 1989

MyxosporeaHenneguya paranaensis Eiras, Pavanelli and Takemoto, 2004

MonogeneaRhinonastes pseudocapsaloideum Kritsky, Thatcher and Boeger, 1988Kritskyia boegeri Takemoto, Lizama and Pavanelli, 2002Tereancistrum curimba Lizama, Takemoto and Pavanelli, 2004Tereancistrum toksonum Lizama, Takemoto and Pavanelli, 2004Tereancistrum sp.Gyrodactylus sp.1Anacanthoroides sp.Ancyrocephalinae

DigeneaSaccocoelioides magnorchis Thatcher, 1978Saccocoelioides nanii Szidat, 1954Unicoelium prochilodorum Thatcher and Dossman, 1975Megacoelium sp.Saccocoelioides leporinodus Thatcher, 1978Saccocoelioides saccodontis Thatcher, 1978Saccocoelioides sp.Tylodelphis sp. (metacercariae)Colocladorchis sp.Sphinterodiplostomum sp. (metacercariae)

Species listLecitobothrioides sp.Digenea (metacercariae)

CestodaProteocephalidea (plerocercoid)Valipora campylancristrota (Wedl, 1855)

NematodaRaphidascaris sp.

AcanthocephalaNeoechinorhynchus curemai Noronha, 1973Quadrigyrus sp.

CopepodaGamidactylus jaraquensis Thatcher and Boeger, 1984Gamispatulus sp.Amplexibranchius sp.Ergasilus sp.

BranchiuraDolops geayi (Bouvier, 1897)

Hirudinea

Erythrinidae• Hoplias aff. malabaricus (Bloch, 1794) (traíra)

DigeneaClinostomum complanatum Rudolphi, 1814 (metacercariae)Prosthenhystera sp.Ithyoclinostomum dimorphum (Diesing, 1850) (metacercariae)Diplostomum sp. (metacercariae)Diplostomum (Tylodelphys) sp.1 (metacercariae)Diplostomum (Tylodelphys) sp.2 (metacercariae)Pseudosellacotyla lutzi (Freitas, 1941)Sphincterodiplostomum musculosum Dubois, 1936 (metacercariae)Diplostomum (Austrodiplostomum) compactum (Lutz, 1928) (metacercariae)Thometrema overstreeti (Brooks, Mayes and Thorson, 1979)

NematodaEustrongylides ignotus Jäegerskiold, 1909 (larvae)Contracaecum sp. (larvae)Porrocaecum sp. (larvae)Paracapillaria piscicola (Travassos, Artigas and Pereira, 1928)Procamallanus (Procamallanus) peraccuratus Pinto, Fábio, Noronha and Rolas, 1976Procamallanus (Spirocamallanus) inopinatus Travassos, Artigas and Pereira, 1928Goezia spinulosa (Diesing, 1839)

AcanthocephalaQuadrigyrus machadoi Fábio, 1983

PentastomidaSebekia oxycephala Sambon, 1922

Isopoda


702 Braz. J. Biol., 69(2, Suppl.): 691-705, 2009


Species list

• Hoplerythrinus unitaeniatus (Spix and Agassiz, 1829) (jejú)

Digenea (metacercariae)NematodaAcanthocephala

Quadrigyrus brasiliensis Machado, 1941

Cynodontidae• Rhaphiodon vulpinus Agassiz, 1829 (dourado-cachorro)

NematodaContracaecum sp. (larvae)


GYMNOTIFORMES

Gymnotidae• Gymnotus carapo Linnaeus, 1758 (morenita)

DigeneaClinostomum complanatum Rudolphi, 1814 (metacercariae)Herpetodiplostomum sp. (metacercariae)Crocodilicola sp.Neodiplostomum sp. (metacercariae)Tylodelphys sp. (metacercariae)

CestodaNomimoscolex chubbi (Pavanelli and Takemoto 1995)

NematodaSpiroxys sp.Hysterothylacium sp. (larvae)Contracaecum sp. (larvae)Eustrongylides sp. (larvae)


• Gymnotus inaequilabiatus (Valenciennes, 1839) (morenita)

DigeneaCestoda

• Gymnotus sylvius Albert and Fernandes-Matioli, 1999 (morenita)

Copepoda

Sternopygidae• Eigenmannia trilineata López and Castello, 1966 (Espadinha)

Copepoda

Rhamphichthyidae• Rhamphichthys hahni (Meinken, 1937) (peixe-espada)

Nematoda

SILURIFORMES

Doradidae• Pterodoras granulosus (Valenciennes, 1821) (abotoado)

DigeneaDadaytrema oxycephala (Diesing, 1836)

Species listCestoda

Monticellia belavistensis Pavanelli, Santos and Takemoto, 1994

NematodaRondonia rondoni Travassos, 1920

Acanthocephala

• Rhinodoras dorbignyi (Kner, 1855) (armado)Nematoda

• Trachydoras paraguayensis (Eigenmann and Ward, 1907) (armadinho)

MonogeneaVancleaveus sp.

DigeneaStrigeoidea (metacercariae)

NematodaProcamallanus sp.Cosmoxynemoides sp.

Auchenipteridae• Auchenipterus osteomystax (Miranda Ribeiro, 1918) (surumanha)

MonogeneaDemidospermus sp.

DigeneaDiplostomum (Austrodiplostomum) compactum (Lutz, 1928) (metacercariae)Creptotrema creptotrema Travassos, Artigas and Pereira, 1928Microrchis oligovitellum Lunaschi, 1987Crocodilicola sp.Clinostomum complanatum Rudolphi, 1814 (metacercariae)Strigeidae (metacercariae)

NematodaProcamallanus (Spirocamallanus) sp.Johnstonmawsonia sp.Rhabdochona acuminata (Molin, 1860)

Copepoda

• Parauchenipterus galeatus (Linnaeus, 1766) (cangati)

MonogeneaDigenea

Microrchis oligovitelum Lunaschi, 1987Clinostomum complanatum Rudolphi, 1814 (metacercariae)

CestodaCangatiella arandasi Pavanelli and Machado dos Santos, 1991

NematodaCucullanellus sp.Goezia sp.

Ageneiosidae• Ageneiosus brevifilis Valenciennes, 1840 (manduvê)

CestodaGibsoniela mandube (Woodland, 1935)


703Braz. J. Biol., 69(2, Suppl.): 691-705, 2009


Species listNematoda

• Ageneiosus ucayalensis Castelnau, 1855 (manduvê)Acanthocephala

Pimelodidae• Pseudoplatystoma corruscans (Spix and Agassiz, 1829) (pintado)

MyxosporeaMonogenea

Pavanelliella pavanellii Kritsky and Boeger, 1998Amphocleithrium paraguayensis Price and Gonzalez Romero 1969Amphocleithrium sp.Vancleaveus fungulus Kritsky, Thatcher and Boeger, 1986Vancleaveus sp.Unibarra sp.Ancyrocephalinae

DigeneaCestoda

Nomimoscolex sudobim Woodland, 1935Nomimoscolex pertierrae Chambrier, Takemoto and Pavanelli, 2006Choanoscolex abscissus (Riggenbach, 1896)Spasskyelina spinulifera (Woodland, 1935)Harriscolex kaparari (Woodland, 1935)Megathylacus travassosi Pavanelli and Rego, 1992

NematodaContracaecum sp.Cucullanus (Cucullanus) pseudoplatystomae Moravec, Kohn and Fernandes, 1993Eustrongylides ignotus Jäegerskiold, 1909 (larvae)Procamallanus (Spirocamallanus) sp.Philometridae

AcanthocephalaBranchiura

Dolops carvalhoi Lemos de Castro, 1949Argulus pestifer Ringuelet, 1948

CopepodaErgasilidaeVaigamidae

• Pimelodus maculatus Lacepède, 1803 (mandi)Monogenea

Demidospermus sp.Unibarra sp.Pavanelliella pavanellii Kritsky and Boeger, 1998

DigeneaThometrema overstreeti (Brooks, Mayes and Thorson, 1979)Crepidostomum platense Szidat, 1954Creptotrema creptotrema Travassos, Artigas and Pereira, 1928Plehniella coelomica Szidat, 1951

Species listProsthenhystera obesa (Diesing,1850)Clinostomum sp. (metacercariae)Diplostomum sp. (metacercariae)

CestodaMonticellia loyolai Pavanelli and Machado dos Santos, 1992Nomimoscolex sp.Valipora sp.

NematodaCucullanus pinnai Travassos, Artigas and Pereira, 1928Spirocamallannus sp.Philometra sp.Monhysterides sp.Goezia sp.Contracaecum sp. (larvae)Eustrongylides sp. (larvae)

AcanthocephalaNeoechinorhynchus sp.Quadrigyrus machadoi Fábio, 1983

CopepodaErgasilus sp.Vaigamus sp.Therodamas sp.Gamidactylus sp.

HirudineaHelobdela sp.Myzobdella sp.

IsopodaTelotha sp.

• Pimelodus ornatus Kner, 1858 (mandi)Cestoda

Mariauxiella pimelodi Chambrier and Rego, 1995Spasskyelina sp.

• Hemisorubim platyrhynchos (Valenciennes, 1840) (jurupoca)

DigeneaCrocodilicola pseudostoma (Willemoes-Suhm, 1870)Sanguinicola platyrhynchi Guidelli, Isaac and Pavanelli, 2002Sphincterodiplostomum sp.

Gorgoderidae

CestodaGoezeella paranaensis Pavanelli and Rego, 1989Spatulifer maringaensis Pavanelli and Rego, 1989

Chambriella itaipuensis (Pavanelli and Rego, 1991)Mariauxiella piscatorum Chambrier and Vaucher, 1999

Nematoda Contracaecum sp. (larvae)


704 Braz. J. Biol., 69(2, Suppl.): 691-705, 2009


Species listCucullanus (Cucullanus) zungaro Vaz and Pereira, 1934Eustrongylides ignotus Jägerskiöld, 1909 (larvae)Goezia sp.


Pentastomida

• Iheringichthys labrosus (Lütken, 1874) (mandi)Monogenea

Demidospermus labrosi França, Isaac, Pavanelli and Takemoto, 2003Demidospermus mandi França, Isaac, Pavanelli and Takemoto, 2003Pseudovancleaveus paranaensis França, Isaac, Pavanelli and Takemoto, 2003

DigeneaHerpetodiplostomum sp. (metacercariae)

CestodaProteocephalidea

NematodaProcamallanus (Spirocamallanus) pimelodus Pinto, Fábio, Noronha and Rolas, 1974Contracaecum sp.

AcanthocephalaQuadrigyrus sp.

• Rhamdia quelen (Quoy and Gaimard, 1824) (bagre)Digenea (metacercariae)

• Pinirampus pirinampu (Spix and Agassiz, 1829) (barbado)

MonogeneaOmothecium sp.Unibarra sp.

Digenea (metacercariae)Cestoda

Nomimoscolex admonticelia (Woodland, 1934)Rudolphiella piranabu (Woodland, 1934)

Nematoda (larvae)Copepoda

Vaigamidae

• Sorubim lima (Bloch and Schneider, 1801) (chinelo)Cestoda

Spatulifer maringaensis Pavanelli and Rego, 1989Paramonticellia itaipuensis Pavanelli and Rego, 1991Nupelia portoriquensis Pavanelli and Rego, 1991Goezeella nupeliensis Pavanelli and Rego, 1991

• Zungaro zungaro (Humboldt, 1821) (jaú)Cestoda

Travassiella avitellina Rego and Pavanelli, 1987

Species listPeltidocotyle rugosa Diesing, 1850Megathylacus brooksi Rego and Pavanelli, 1985Jauella glandicephalus Rego and Pavanelli, 1985Chambriella agostinhoi Pavanelli and Santos, 1992Choanoscolex abscissus (Riggenbach, 1895)

NematodaCucullanus sp.

Hypophthalmidae• Hypophthalmus edentatus Spix and Agassiz, 1829 (mapará)

Nematoda

Callichthyidae• Hoplosternum littorale (Hancock, 1828) (tamboatá)

DigeneaClinostomum complanatum Rudolphi, 1814 (metacercariae)

Loricariidae• Loricariichthys platymetopon Isbrücker and Nijssen, 1979 (cascudo-chinelo)

MonogeneaDigenea

Clinostomum complanatum Rudolphi, 1814 (metacercariae)Crocodilicola pseudostoma (Willemoes-Suhm, 1870) (metacercariae)

NematodaRaphidascaris (Sprentascaris) mahnerti (Petter and Cassone, 1984)

• Loricariichthys rostratus Reis and Pereira, 2000 (cascudo-chinelo)

Digenea (metacercariae)• Hypostomus regani (Ihering, 1905) (cascudo-chita)

DigeneaDiplostomum (Austrodiplostomum) compactum (Lutz, 1928) (metacercariae)

• Rhinelepis aspera Spix and Agassiz, 1829 (cascudo-preto)

ProtozoaTrypanosoma nupelianus Eiras, Rego and Pavanelli, 1990

• Megalancistrus parananus (Peters, 1881) (cascudo-abacaxi)

ProtozoaTrypanosoma guairaensis Eiras, Rego and Pavanelli, 1989

Hirudinea

SYNBRANCHIFORMESSynbranchidae

• Synbranchus marmoratus Bloch, 1795 (mussum)Digenea


705Braz. J. Biol., 69(2, Suppl.): 691-705, 2009


Species list

PERCIFORMES

Sciaenidae• Plagioscion squamosissimus (Heckel, 1840) (curvina)

MonogeneaDiplectanum piscinarius Kritsky and Thatcher, 1984

DigeneaDiplostomum (Austrodiplostomum) compactum (Lutz, 1928) (metacercariae)

Nematoda (larvae)

Cichlidae• Geophagus proximus (Castelnau, 1855) (acará)

MonogeneaSciadicleithrum sp.1Sciadicleithrum sp.2

DigeneaAscocotyle sp. (metacercariae)

NematodaRaphidascaris (Sprentascaris) sp.

Acarina

• Satanoperca pappaterra (Heckel, 1840) (acará)Monogenea

Cleidodiscus sp.Sciadicleithrum sp.

DigeneaDiplostomum (Austrodiplostomum) compactum (Lutz, 1928) (metacercariae)Ascocotyle sp. (metacercariae)


NematodaCopepoda

• Cichla kelberi Kullander and Ferreira, 2006 (tucunaré)Monogenea

Gussevia arilla Kritsky, Thatcher and Boeger, 1986Gussevia longihaptor (Mizelle and Kritsky, 1969)Gussevia undulata Kritsky, Thatcher and Boeger, 1986

DigeneaClinostomum sp. (metacercariae)

Species listDiplostomum (Austrodiplostomum) compactum (Lutz, 1928) (metacercariae)Diplostomum sp. (metacercariae)

CestodaProteocephalus macrophalus (Diesing, 1850)Proteocephalus microscopicus (Woodland, 1935)Sciadocephalus megalodiscus Diesing, 1850

NematodaContracaecum sp. (larvae)

AcanthocephalaQuadrigyrus machadoi Fábio, 1983 (larvae)

Isopoda

• Astronotus crassipinnis (Heckel, 1840) (oscar)Monogenea

Gussevia asota Kritsky, Thatcher and Boeger, 1989Gussevia astronoti Kritsky, Thatcher and Boeger, 1989Gussevia rogersi Kritsky, Thatcher and Boeger, 1989Gussevia sp.

• Crenicichla britskii Kullander, 1982 (joaninha)Monogenea

Sciadicleithrum sp.DigeneaCestoda

Valipora sp.Nematoda

• Crenicichla niederleinii (Holmberg, 1891) (joaninha)Monogenea

Sciadicleithrum sp.DigeneaNematoda

• Cichlasoma paranaense Kullander, 1983 (cará)Acanthocephala

• Laetacara sp. (cará)

MonogeneaCleidodiscus sp.

Digenea

Documents

Diversity of parasites of fish from the Upper Paraná River ... · zoa, three species of Phylum Sarcomastigophora were reported: Trypanosoma guairaensis Eiras, Rego and Pavanelli,